Life Sciences Contribution O7 Royal Ontario Museum New Systematic Data for the North American Caddisfly Genera Lepania, Goeracea and Goerita (Trichoptera: Limnephilidae) Glenn B. Wiggins RC ; é M http:/;www.archive.org/details/newsystemati na ee an Tee, 4 ees PP Rak 4, We Lia Py, by dy ’ ay »* + . ns he) A iol | : a) >< 4 “aA 4 \ =< ’ 4 rt ; _ i ‘ Ar a f yy 4 4 ? f le - i ‘ o hf “2 J 5 5 . : > Whaat 4 = wa r. he re » ¥ 4 w Fs tt rs = - 7 K ee , ™ 4 * rT *~ . 4 * ‘ — q ‘ re ; A “; ‘ P * _ 1S i i : 7 : i es ; ; be, Ue + e 1% : ma _ waht a ' ro art > ~ 4 J , - ¥ ’ “ 7 < ¥ > J MY : ‘ x { = i i a] e : hs o e fics ‘ 4 "i In 2011 with fundies from Ray / \ os + ‘i 5 a ‘ ; a. . - ite 1. 2 d . 4204 = + - j t 3 +a] 7 ai y nive Toron’ | ~ " — a ¢ ‘ ‘ a z Car ape oe ae 4 PX * 5 = - . 4‘ ‘ “- = x i #) P - = ie G . ' u :! re = a a ; “ f y ‘ ? . rn = & ' " bas = 4 5 7 4 ms r . a - al 4 > , 4, Hi, A 7 : i eo es ‘ : a . . * vy va y) : : ia i = 7 a 4 s = : . t - 3 ~~ ) , ai aan + he Pe * ‘ cus ‘ : . } . - é = hey ; fp F oF. ‘ e " rH , 2 y BS a 7 mt; re - a 6 ae Avie - . sr ' f ra Pts Sie ¢ ; | a . L . " : = ~ P 7 ‘ ‘ft < +. Lf oe » ie ' r Ls aren ie Foy ~_ ” 1 . = . d = ivf a4 ee LIFE SCIENCES CONTRIBUTIONS ROYAL ONTARIO MUSEUM NUMBER Q] GLENN B. wicciIns New Systematic Data for the North American Caddisfly Genera Lepania, Goeracea and Goerita (Trichoptera: Limnephilidae) Publication date: 27 June 1973 Suggested citation: Life Sci. Contr., R. Ont. Mus. ROYAL ONTARIO MUSEUM PUBLICATIONS IN LIFE SCIENCES The Royal Ontario Museum publishes three series in the Life Sciences: LIFE SCIENCES CONTRIBUTIONS, a numbered series of original scientific publi- cations, including monographic works. LIFE SCIENCES OCCASIONAL PAPERS, a numbered series of original scientific publications, primarily short and usually of taxonomic significance. LIFE SCIENCES MISCELLANEOUS PUBLICATIONS, an unnumbered series of publications of varied subject matter and format. All manuscripts considered for publication are subject to the scrutiny and editorial policies of the Life Sciences Editorial Board, and to review by persons outside the Museum staff who are authorities in the particular field involved. LIFE SCIENCES EDITORIAL BOARD Chairman, R. L. PETERSON Editor, J. R. TAMSITT Associate Editor, D. BARR Associate Editor, E. J. CROSSMAN GLENN B. WIGGINS is Curator of the Department of Entomology and Invertebrate Zoology, Royal Ontario Museum, and Professor in the Department of Zoology, University of Toronto. PRICE: $2.50 ©The Royal Ontario Museum, 1973 100 Queen’s Park, Toronto, Canada PRINTED BY MaCKINNON-MONCUR New Systematic Data for the North American Caddisfly Genera Lepania, Goeracea and Goerita (Trichoptera: Limnephilidae) Abstract In a brief synopsis of the North American genera Lepania, Goeracea, and Goerita, larvae and pupae are described for the first time for Lepania cascada Ross, Goeracea genota (Ross), Goeracea oregona Denning, and Goerita betteni Ross. Keys to larvae, males, and females are given for the two species now known in each of the genera Goeracea and Goerita. For Lepania cascada an analysis is made of systematic characters of larvae, pupae and adults, demonstrating the probability that true affinities of the species lie with the limne- philid subfamily Goerinae rather than with the Apataniinae, to which it has been most recently assigned. A new tribe, Lepaniini, is created in the Goerinae for the genus. Based on retention of several primitive characters, Lepania is regarded as a genus of unusual phylogenetic significance, annectent between the ancestral stock of the Limnephilidae (auct., excluding Goeridae) and the Goerinae, treated in this paper as a subfamily of the Limnephilidae. Introduction The purpose of this paper is to present new information on larval and pupal _ Stages of species in the North American caddisfly genera Lepania, Goeracea, and Goerita. A synopsis of each genus is offered, including keys to species for larvae, males and females, where known, in Goeracea and Goerita. Made known for the first time are larvae and pupae of Lepania cascada, Goeracea genota, G. oregona, and Goerita betteni. Assessment of the systematic data made available for Lepania cascada establishes that genus as one particularly significant in interpreting evolu- tionary lines within the family Limnephilidae. Furthermore, there are implications for two areas of the systematics of Trichoptera in general. Firstly, further reinforcement is provided for the view (Ross, 1956; Wiggins, 1964) that knowledge of larval stages is critical in interpreting systematic relationships of family-group taxa. Secondly, the taxonomic gap between the Limnephilidae and Goeridae (auct.) is reduced, pointing up again the lack of agreement among students of caddisflies concerning recognition of the Goeridae as a distinct family or as a subfamily of the Limnephilidae. The latter course is adopted here. Materials and Methods Collections and observations upon which this paper is based were made during a series of field expeditions to western and southeastern North America carried out by staff of the Department of Entomology and Inverte- brate Zoology, ROM. Unless indicated otherwise, all specimens are deposited in the Royal Ontario Museum. Numbers in parentheses cited in Material Examined refer to field collection data sheets of the Department of Ento- mology, ROM. Associations between larval, pupal, and adult stages were established through collection of larvae and mature pupae or by collection of larvae and adults from the same site. Measurements were made with a Zeiss eyepiece micrometer at 10X magnification, and converted to milli- metres by a Leitz calibration scale. Quantitative data, expressed as a range from minimum to maximum, are followed by the number of specimens, in parentheses, on which the figures are based. Length of larva expresses the length of the linear axis between anterior and posterior limits of larvae in the position shown in the habit illustrations (Figs. la, 10a, 17a). Systematic Data Lepania Lepania Ross, 1941, p. 102; type species L. cascada Ross. Lepania cascada Ross Lepania cascada Ross, 1941, p. 102, fig. 81. Lepania cascada. Schmid, 1955, p. 69, figs. 43, 44. Lepania cascada. Schmid, 1968a, p. 685, figs. 23-27. The single species assigned to this genus is recorded from a few localities in Oregon and Washington. L. cascada has been known for 30 years but until now only in the adult stage. When first described from the adult (Ross, 1941), it was assigned simply to the Limnephilidae but was later placed in the subfamily Discosmoecinae (Schmid, 1955). After a re- evaluation of the morphology of the adults, Lepania was later assigned to the Apataniinae (tribe Moropsychini) by Schmid (1968a). Recent dis- covery of larvae, mature pupae, and adults in Oregon has provided the first associated series for study. LARVA (Figs. 1-3, 8-9). Larvae of L. cascada have a general resemblance to those of the Goerinae, with which they share an elongate mesepisternum and subdivided mesonotal plates; they can be distinguished most readily from genera in that group by the presence of single gills and by a distinct transverse division in the mesal pair of mesonotal sclerites; other diagnostic characters are outlined below. Figs. 1-4 Lepania cascada larva. 1. Larva: a, lateral view, X26; b, mesotarsal claw, X200; c, ninth abdominal segment and anal prolegs, dorsal view, X60; d, claw of right anal proleg, lateral view, X290. 2. Head and thorax: a, dorsal view, X55; b, head capsule ventral view, X55; c, right side of head, dorsal view, X55. 3. Mandibles, ventral view X150. 4. Larval case: a, lateral view, X20; b, posterior end of case, X159. N \ SS ~ : S EX\. ICEL fl fA ANNE \\\\\ Ns ig Ye\ it (Pl f | SS lateral YS -sclerite GU jp, i, Xi SAN § 2G) ~—basal tuft Ic Length of final-instar larva, 4.6-5.3 mm (5). All sclerotized parts reddish brown in colour. Head (Fig. 2a) circular in dorsal view, surface finely pebbled, dorsum with many secondary setae; each antenna originating in a depression (Fig. 2c); shape of gular sclerite (Fig. 2b) an elongate triangle with concave lateral margins. Labrum (Fig. 9) with anterolateral edges unpigmented and semimembranous. Mandibles (Fig. 3) with cutting edges retaining individual pointed teeth; a thick brush of feather-like hairs arises from mesal edge of each mandible. Labium (Fig. 8) with two, quadrate, submental sclerites, each unusual in possession of two large setae in place of the usual one; sclerite of palpiger angulate but not continued around base of labial palp. Maxilla (Fig. 8) unusual in having about six stout setae along lateral edge of each sclerite of stipes; sclerite of palpifer with small, separate, mesal piece; galea low and rounded. Pronotum (Fig. 2a) relatively large, longer than wide, anterior margin straight, lateral and posterior margins evenly confluent along a prominent ridge; median portion of pronotum arched dorsad; prosternal horn well developed. Mesonotum with three separate sclerites, possibly corresponding to sa 1, sa 2 and sa 3; mesopleuron strongly developed, mesepisternum extended anterad as a stout pointed process, concave dorsally. Metanotum with all three primary sclerites well developed but separate, sa J sclerites adjacent on the midline. Legs (Fig. la) with trochanteral brushes absent; femora of second and third legs with several stout setae along ventral edge, basal seta of tarsal claws much shorter than claw (Fig. 1b). Abdomen (Fig. la) with humps of segment I well developed, approxi- mately 31-43 small setae between the median hump and each lateral hump, approximately 106-128 setae on venter of segment I (5 specimens, final instar); lateral humps with minute spines. Gills single, present only on segment UI, which bears a posterodorsal and posteroventral pair, each gill with a swollen base. Venter v and vi each with an ovoid, sclerotized ring. Lateral line of setae absent. Dorsal sclerite of segment Ix (Fig. Ic) with two pairs of large setae, 33-39 smaller setae (5, final instar). Anal prolegs (Fig. 1c) with numerous, small setae on each lateral sclerite, basal tuft with two setae on each side much larger than the others, sinuate near bases; small sclerite ventrolaterad of each basal tuft; claw of anal proleg lacking accessory tooth (Fig. 1d). Larval case (Fig. 4) of small rock pieces, strongly tapered, slightly curved, pieces on ventral surface smaller than those on dorsolateral; posterior opening restricted with silk into an angulate slit that in profile shows dorsal lip to overhang ventral. Length of final-instar larval case, 4.8-5.3 mm (5). Figs. 5-9 Lepania cascada. 5. Head of pupa, facial view, X60. 6. Pupal case: a, anterior closure membrane, X19; b, posterior closure membrane, facial view, X19; c, same as b, in profile X19. 7. Abdomen of pupa: a, abdomen, X23, with sclerotized plates enlarged, dorsal view; b, anal processes, dorsal view, X84. 8. Larva: a, maxillae and labium ventral view, X210; b, maxilla from right side of a, enlarged, ventral view, X428. 9. Labrum of larva, dorsal view, X159. fii MN) wo ly, & = 7, 5) om 6) 8b af (: 7) ww \ \ Y YY Yj Uy Yy PUPA (Figs. 5-7). Pupae of L. cascada can be recognized by the curved dentate apex of the anal processes. Length of pupa, 5.6-6.5 mm (6). Head (Fig. 5) with mandibles finely toothed, cutting edge convex proximal to apex as in Apatania; labrum with five pairs of stout setae, all strongly hooked apically, a group of five, small setae on each side at base of labrum; pair of setae in middle of face differing from Apatania and allies in being much longer and hooked apically. Abdomen (Fig. 7) with sclerotized plates as illustrated, not greatly different from Apatania and allies; gills absent; anal processes long, slender, each curving smoothly laterad well before apex, lateral face of curve dentate, three stout setae evenly spaced on mesal surface of curve. Pupal case with anterior silken membrane (Fig. 6a) incorporating a few small stones, small openings around periphery; posterior membrane (Figs. 6b, c) basically as in larval case, but with further restriction of angulate opening with silk, and with dorsal lip of opening still overhanging ventral lip. Length of pupal case, 6.6-7.2 mm (3). ADULT. Adults of L. cascada can be distinguished by characters of the male and female genitalia, which are similar to no other species. Length of forewing (both sexes), 5.9-6.6 mm (15). Overall colour black with white fringes on both wings. Head with one pair of warts on dorsum posterior to median ocellus (Fig. 22) and one posterolateral pair behind lateral ocelli; maxillary palpi (Fig. 23) of male curved dorsad before face, with three segments, terminal segment not thickened, approximately as long as basal and middle segments combined. Mesoscutum (Fig. 22) with a pair of small, elongate warts; mesoscutellum with a single ovoid wart. Spurs 1, 2, 3. Wing venation (Fig. 21) similar in the sexes, R, of hind wings incomplete; wing-coupling mechanism (Fig. 21) well developed, consisting of a row of stout, hooked setae along costal margin of hind wing, and a short band of very long setae arising from trailing edge of forewing, closely appressed to it. Male genitalia (Figs. 25, 26). Segment 1x wide laterally, narrower dorsally and ventrally; membrane of segment VIII attached along a line posterior to anterolateral edge; both dorsum and venter of 1x with conspicuous semi- membranous areas, delimited laterally by sclerotized ridges. Claspers two- segmented, basal segment short laterally, wide caudally; terminal segment flat, blade-like, strongly curved. Segment x consisting of four pairs of processes (Fig. 26): large praeanal appendages, each with a short mesal branch in dorsal view; ventral to these a pair of short, finger-like lobes (here designated accessory appendages); ventral to these arises a short plate, apparently formed by fusion of two heavily-sclerotized, blade-like processes, upturned apically (internal branches); from base of preceding processes arises a pair of slender, finger-like processes curved dorsad (external branches). Phallus with invaginated endotheca bearing two pairs of stout, sclerotized processes, dorsal pair with bases united into pointed, median prong. Female genitalia (Fig. 30). Sternum of segment vill with an unusual median division, divided plate free for about half its length from membranous body 6 wall along its posterior margin; on each side is a large sclerite. Dorsum Ix not evident (either fused completely with x or absent); venter Ix with a pair of lateral sclerites forming two stout sclerotized lobes in ventral view, and between these lies a lightly sclerotized area with a median apical notch bearing a pointed, tongue-like process within it. Segment x (including dorsum Ix?) fused with supragenital plate to form a broad, flattened chamber enclosing anal opening. MATERIAL STUDIED. OREGON. Benton Co.: Marys Peak, seepage area beside Parker Creek at Marys Peak Campground, 14 June 1967, 3d6, 4?9; same data as above, 14 and 15 June 1968, 66, 2°, larvae; same data as above, 6 June 1969, (690050) 6d, 2°, pupae, larvae. WASHINGTON: Mount Rainier National Park, small spring run, 12.1 mi E of Longmire on Route 706, 4 July 1969, (690168) 1 larva. All collected by Rom field parties. HABITAT. Larvae and mature pupae of L. cascada taken on Marys Peak, Oregon, were collected from the muck of a spring seepage area several feet from a small spring stream, Parker Creek, but not in the stream. Spring seepage emerged from the ground at the top of an inclined bank covered with a dense growth of moss and Equisetum. Water overflowed from small depressions in the soil, dripped progressively to lower levels, and ultimately joined the stream. Soil in the whole area was saturated with water. Adults were flying actively in the rich, herbaceous plant growth over the seepage area. Adults, larvae, and pupae of L. cascada and of Moselyana comosa were collected from this site at the same time. Goeracea Goeracea Denning, 1968, p. 24; type species Goerita genota Ross. This genus comprises two species, G. genota (Ross) and G. oregona Denning, both confined to western North America. Larvae and pupae were not known previously for either species. LARVA (Figs. 10-11, 16). Larvae of this genus can be distinguished from other North American goerine larvae by the presence of single gills and by the dorsally-flattened, subcircular pronotum. When wholly withdrawn into its case, the larva can effectively seal the anterior opening with its pronotum, the part of the mesonotum anterad of the transverse ridge, and the mesepis- terna. Pressed together these parts form a circular “operculum,” with the head curled ventrad beneath the pronotum. Length of larva to 6.6 mm. All sclerotized parts reddish-brown. Head (Fig. 11a) ovoid, rather small in relation to pronotum, inserted well below dorsum of pronotum, and strongly inclined ventrad when larva is withdrawn into case; dorsum finely pebbled, lacking secondary setae, bases of primary setae nos. 14 and 15 arising close together; each antenna located in a shallow concavity; gular sclerite (Fig. 11b) acorn-shaped; labrum (Fig. 11d) with conspicuous anterolateral membranous border; mandibles (Fig. Fé llc) with smooth cutting edges, teeth absent, hairs of mesal brush rough- ened along their length; labium (Fig. 16) with a pair of irregularly-shaped submental sclerites, each with one stout seta; sclerite of palpiger straight, not curved around base of labial palp; maxilla with only one lateral seta on each stipital sclerite, galea flat. Pronotum (Fig. 11a) massive, dorsum flat and subcircular; anterior and lateral edges of pronoium very thick; prosternal horn prominent. Mesonotum with sclerites contiguous, mesal pair of sclerites undivided; mesepisternum compressed, very long and stout; a sharp ridge extends transversely across mesonotum, at which head and thorax are canted forward at a prominent angle to remainder of thorax and abdomen. Legs (Fig. 10a) lacking tro- chanteral brushes, femoral setae sparse and weakly developed; basal seta of tarsal claws long, reaching to tip of claw (Fig. 10b). Abdomen (Fig. 10a) with humps of segment 1 well developed, approxi- mately 42-58 setae between median hump and each lateral hump, approxi- mately 139-174 setae on ventral surface of first segment. Gills singie. Venters of segments Iv, v, and vi each with an ovoid sclerotized ring. Lateral line of setae absent. Dorsal sclerite of segment Ix with approximately 27-47 setae, only a mesal pair much stouter than the others. Anal prolegs (Fig. 10c) with only one or two setae on lateral sclerite, basal tuft with two setae, very stout but rather short and sinuate, their bases surrounded by a small sclerite; claw of anal prolegs without an accessory tooth (Fig. 10d). Larval case (Fig. 12) of small, rock pieces, with a row of larger pieces along each side; posterior opening restricted with silk except for a small circular hole dorsal to centre. Length of larval case to 6.1 mm. PUPA (Figs. 13, 14). Similar to Goerita. Length 5.0-6.4 mm. Head (Fig. 13) with cutting edges of mandibles convex proximal to apex, lacking fine teeth; labrum with five pairs of stout setae, all strongly hooked apically, a group of three setae on each side at base of labrum. Abdomen (Fig. 14a) with sclerotized plates as illustrated, without distinctive features; gills absent; anal processes (Figs. 14b, c) very short and stout, heavily sclerotized, apices curved dorsad and bluntly pointed; three short setae arising near apex of each process. Pupal case with anterior closure membrane (Fig. 15) usually incor- porating one relatively large stone for about one-half of its area; ventral to this stone is a meshed opening in the silk through which labral bristles protrude; posterior opening apparently unchanged from small, eccentric hole in larval case. Length of pupal case 5.0-7.5 mm. Figs. 10-12 Goeracea genota larva. 10. Larva: a, lateral view, X24; b, mesotarsal claw, X290; c, ninth abdominal segment and anal prolegs, dorsal view, X64; d, claw of right anal proleg, lateral view, X272. 11. Head and thorax: a, dorsal view, X39; b, head capsule, ventral view, X39; c, right mandible, dorsal view, X92; d, labrum, dorsal view, X119. 12. Larval case: a, oblique dorsal view, X 15.6; b, posterior end, X15.6. x \ 5 ANGE: WS: \' SS Ulli ADULT. Apart from genitalic characters, Goeracea can be distinguished from Goerita by the narrower posterodorsal warts of the head and by the unmodi- fied maxillary palpi of the male. Goeracea can be distinguished from Lepania by a well-developed R, in the hind wing. Length of forewing (both sexes), 4.3-8.0 mm. Colour dark brown. Head with dorsal warts much as in L. cascada (Fig. 22), posterior warts narrower than in Goerita; maxillary palpi of male with terminal and middle segments about same length; all segments normal in thickness (Fig. 23). Thorax as in L. cascada (Fig. 22). Forewings of male with coarse scales on dorsal surface along Sc, R2, and R; and in anal area of wing; frenulum consisting of four or five stout setae. Venation similar in the sexes, much as in Goerita (Ross, 1938, fig. 113), but with discoidal cell closed by a crossvein. Spurs 2, 3-4, 4. Male genitalia (Figs. 28, 29). Primarily distinctive in the structure of segment x, where the praeanal appendages (interpreted here as in Fig. 28b) are widely separated by the membranous body of that segment; other appendages of xX apparently lacking. Claspers much as in Goerita; basal segment scoop-shaped, with a prominent ventromesal process; terminal segment finger-like. Phallus with invaginated endotheca bearing two slender spines and two stout double-pointed spines. Female genitalia (Figs. 32, 33). Tergum of segment Ix evidently represented by a small sclerite separated from what appears to be tergum x (here inter- preted as in Fig. 32a). Venter 1x with a pair of lateral sclerites extending across posterior margin, these separated on meson by a triangular, median plate. HABITAT. Both species occur in small, cold, spring streams, where larvae are usually found on rocks. Figs. 13-16 Goeracea genota. 13. Head of pupa, facial view, X52.7. 14. Pupa: a, ab- domen, X28, with sclerotized plates enlarged; b, anal processes, dorsal view, X94.7; c, same as in b, lateral view, X94.7. 15. Anterior closure membrane of pupal case, X15. 16. Maxillae and labium of larva, ventral view, X228. 10 2 Key to species of Goeracea . Posterodorsal and posteroventral gills present only on abdominal segment I (Fig. 10a), occasionally with posterodorsal gills on Iv; metanotal sa / represented by 1 or 2 setae, usually without a basal sclerite (Fig. 11a); length to 6.6 mm Posterodorsal and posteroventral gills present on abdominal segments II and Iv, posterodorsal gills on segment v; metanotal sa / usually represented by a small setose sclerite; length to 5.4 mm Males... Jf 04. Female... Ss Se eS. . Basal segment of clasper with ventromesal process broad and tapering abruptly in ventral view (Fig. 29b; Ross, 1941, fig. 93); segment x with elon- gate, praeanal appendage along each side, bear- ing a narrow, thumb-like lateral lobe (Fig. 29a)... Basal segment of clasper with ventromesal process slender and tapering gradually to an acute apex in ventral view (Fig. 28c); praeanal appendages of segment x with lateral lobe broad and triangular (Riga Sb). ot eee ae ee ae. Se ae . Median plate of venter Ix sclerotized and pigmented, thus largely delimited from remainder of segment in ventral view (Fig. 33a); posteromedian notch of segment x uniformly broad (Fic: 33b) 2 272... Median plate of venter 1x little sclerotized and pigmented, thus barely delimited from remainder of segment in ventral view (Fig. 32b); postero- median notch of segment x narrow and with a G. oregona en _ G. genota Constaction® (Fig: 32C) (52. 2 eee neem ce G. oregona Goeracea genota (Ross) Goerita genota Ross, 1941, p. 116, fig. 93. Goeracea genota. Denning, 1968, p. 24, figs. 12, 13. Insects of this species are generally slightly larger than G. oregona in corresponding larval, pupal, and adult stages, and have more setae on abdominal segment I of the larva, but fewer on the dorsal sclerite of Ix. LARVA (Figs. 10-12, 16). Length of final-instar larva 5.3-6.6 mm (25); length of larval case, same specimens 5.0-6.1 mm. Abdominal segments (final instar): I, setae between median hump and each lateral hump 51- 58 (5); setae on ventral surface 156-174 (5); Ix, setae on dorsal sclerite 27-34 (5). PUPA (Figs. 13-15). Length of pupa 5.6-6.4 mm (5); length of pupal case, same specimens 7.0-7.5 mm. ADULT (Figs. 29, 33). Length of forewing (both sexes), 6.9-8.0 mm (5). BIOLOGY. Records indicate that adults of this species generally emerge earlier (April) than do those of G. oregona. A life cycle of 2 years is demonstrated by emergence of an adult in the laboratory in July 1969 from a series of final-instar (= n) larvae collected in June 1968. The 2-year cycle is further borne out by the number of different instars available at a given time in our collections (all stages from n-2 larvae to adults in April). Larvae of this species occur in cool streams, sometimes in a film of water flowing over rocks and logs. DISTRIBUTION. This species is known from British Columbia, Idaho (Smith, 1965), Oregon, and Washington. MATERIAL STUDIED. BRITISH COLUMBIA: Cultus Lake, small streams on east side, between Entrance Bay and Maple Grove Campgrounds, 20 June 1969, RoM Field Party (690113), 1 larva; Port Renfrew, small spring runs crossed by road, 15.9 mi W Shawnigan Lake, 28 June 1969, Rom Field Party (690138), 10 larvae; Victoria, small spring run entering Goldstream River at Goldstream Campground, 28 June 1969, Rom Field Party (690142), 1 larva. IDAHO. Valley Co.: North Fork Payette River and Canyon Forest Camp Creek, 8.5 mi N Banks on Highway 15, 13 April 1964, S. D. Smith (O01-SDS), 864, 1°; North Fork Payette River, 9 mi S Smith Ferry, 14 April 1965, S. D. Smith and E. R. Logan, 6d¢6. (All Idaho specimens in S.D. Smith collection. ) OREGON. Benton Co.: Alder Creek, 7 mi E Alsea on Route 34, 4-15 April 1964, G. B. Wiggins, R. S. Scott, 5 larvae, 2dd; small spring, 7 mi E Alsea on Route 34, 4 April 1964, G.B.W., R.S.S., 2dd, 1° ; Marys Peak, 12 April 1964, G.B.W., R.S.S., 4 larvae; Marys Peak, Chintimini Creek, 3-14 April 1964, G.B.W., R.S.S., 13 larvae, 3 pupae, 366; Marys Peak, ig Chintimini Creek, 25 Sept. 1966, Rom Field Party, 1 larva; Marys Peak, spring runs near Chintimini Creek, 5 July 1963, G.B.W., 1 larva; Marys Peak, North Fork Rock Creek, 5 July 1963, G.B. W., 7 larvae; Marys Peak, North Fork Rock Creek, 29 Aug. 1963, N.H. Anderson, 1 larva; Marys Peak, North Fork Rock Creek, 13 April 1964, G.B.W., R.S.S., many larvae, 8 pupae; Marys Peak, Road 1296, 0.6 mi off Marys Peak Road, stream no. 1, sample no. 1, 2 July 1968, J. Wold, 14 larvae; Marys Peak, Road 1296, 0.5 mi off Marys Peak Road, stream no. 0, 11 July 1968, J. Wold, 9 larvae; Marys Peak, Rock Creek Road, small streams, 14 April 1964, G.B.W., R.S.S., 36d, 12° ; Marys Peak, spring run entering Woods Creek, 6 April 1964, G.B.W., R.S.S., 2 larvae:' Hood: River Co::.31-March: 1962,8:G. Jewett Ir., cdc. Josephine Co.: Oregon Caves National Monument, Cave Creek, 100 yds below Chateau, elevation 3800 feet, 14 June 1963, N.H. Anderson, 2 larvae; No Name Creek, near Oregon Caves National Monument, elevation 3200 ft, 14 June 1963, N.H. Anderson, 1 larva. Lincoln Co.: Slide Camp- ground, Route 34 E Tidewater, 15 April 1964, G.B.W:, RSS., f9. Linn Co.: Quartzville Road, small runs approximately 1.6 mi E Yellow- stone Guard Station, 16-22 June 1968, rom Field Party, many larvae, 1 pupa; Quartzville Road, small stream 1.7 mi W Yellowstone Guard Station, 22 June 1968, Rom Field Party, 15 larvae, 16 (reared); Quartzville Road, small stream 2.5 mi W Yellowstone Guard Station, 16-17 June 1968, ROM Field Party, 18 larvae. Multnomah Co.: Corbett, Crown Point, spring run, 15 July 1963, G.B.W., 15 larvae; Corbett, Crown Point, spring run, 17 April 1964, G.B.W., R.S.S., 5 larvae. WASHINGTON. Pierce Co.: Mount Rainier National Park, small runs on rock cut, 1.6 mi S Crystal Mountain Road, Route 410, 16 June 1969, Rom Field Party (690086), many larvae; Mount Rainier National Park, Long- mire, small streams entering Fish Creek near Westside Road, 3 July 1969, ROM Field Party (690164), 5 larvae; Mount Rainier National Park, Long- mire, seepage stream crossed by Westside Road, elevation 2500 ft, 13 June 1969, Rom Field Party (690074), 13 larvae. Jefferson Co.: Olympic National Park, small streams crossing Hoh Valley Trail, between Hoh River Campground and Tone Creek Junction, 30 June 1969, Rom Field Party (690158), 17 larvae. G oeracea oregona Denning Goeracea oregona Denning, 1968, p. 25, figs. 14, 15. Insects of this species are generally slightly smaller than G. genota in corresponding stages, and have fewer setae on abdominal segment | of the larva, but more on the dorsal sclerite of Ix. LARVA. Length of final-instar larva 4.0-5.4 mm (25); length of larval case, same specimens, 3.9-5.0 mm. Abdominal segments (final instar): I, setae between median hump and each lateral hump 42-51 (5); setae on ventral surface 139-158 (5); 1x, setae on dorsal sclerite 36-47 (5). A single larva 14 from Mt. Tamalpais near San Francisco, California, has the gill complement of G. oregona, but metanotal sa J is represented by two setae without a sclerite, a condition found to be concordant with the restriction of gills to segment I, evidently typical of G. genota in the material studied. No adults from this site are available, and the larva is a penultimate instar. The specimen is tentatively assigned to G. oregona. PUPA. Length of pupa 5.0-5.8 mm (4); length of pupal case 6.6-8.2 mm (10). In this species the anal processes tend to be more sharply curved dorsad than in G. genota (Fig. 14). ADULT. Length of forewing (both sexes), 4.3-4.9 mm (5). BIOLOGY. The California records indicate that adults of this species tend to emerge somewhat later (June to August) than those of G. genota. The Oregon record for adults of G. oregona on May 22 (Denning, 1968) appears to bridge much of this time gap between the two. Collection of pupae, final instar (n) larvae, n-2, and n-3 instar larvae in July suggests that 2 years are required for completion of the life cycle. Collection of pupae and n-2 instar larvae in October is noteworthy. DISTRIBUTION. Known previously only from Jackson County in south- western Oregon (Denning, 1968), the range of this species is extended to include the Sierra Nevada and perhaps the Coast Range of California. MATERIAL STUDIED. CALIFORNIA. Marin Co.: Mount Tamalpais State Park, small stream, 13 Oct. 1966, Rom Field Party, 1 larva. Nevada Co.: Boca Spring, 6 mi NW of Truckee, elevation 5900 ft, 6 June 1966, G.B. W., A.L. Sheldon, many larvae, 26d; Univ. California Sagehen Creek Research Project, 8 mi NW of Truckee, Kiln Meadow Spring, tributary Sagehen Creek, elevation 6500 ft, 19 June 1966, G.B.W., A.L. Sheldon, many larvae, many pupae, 36d, 49°; Univ. California Sagehen Creek Research Project, 8 mi NW of Truckee, Kiln Meadow Spring, tributary Sagehen Creek, elevation 6500 ft, 27 July 1966, A.L. Sheldon, 1¢; Univ. California Sagehen Creek Research Project, 8 mi NW of Truckee, Kiln Meadow Spring, tributary Sagehen Creek, elevation 6500 ft, 7 Oct. 1966, ROM Field Party, many larvae, many pupae; Univ. California Sagehen Creek Research Project, 8 mi NW of Truckee, spring stream, tributary Sagehen Creek, 20 Aug. 1966, A.L. Sheldon, 1d. Goerita Goerita Ross, 1938, p. 171; type species Goerita semata Ross. As a consequence of assignment of the western species originally treated under this name to the genus Goeracea, Goerita now comprises but two species. Both species occur in small, spring streams and are known only from the eastern United States. They are exceedingly local in distribution. |) LARVA (Figs. 17, 18, 20). Larvae of this genus can be distinguished from all other North American goerine larvae by the absence of abdominal gills and by the prominent median ridge on the pronotum. Length of larvae to 6.1 mm. All sclerotized parts reddish brown, those of head and thorax pebbled. Head (Figs. 18, 20) with indistinct median and dorsolateral ridges, otherwise structures of head as in Goeracea. Pronotum (Figs. 18, 20) strongly arched dorsomesally, posterior two- thirds with a raised ridge on each side of midline; anterior margin with prominent anterolateral point on each side; prosternal horn prominent. Mesonotum with mesal pair of sclerites undivided, an angulate ridge extending transversely across mid part of segment; mesepisternum com- pressed, not as long as in Goeracea. Metanotum with all three primary setal areas represented by setose sclerites, sa / sclerites broadly separated on the meson. Legs much as in Goeracea. Abdomen with humps of segment I well developed, about 20-58 setae between median hump and each lateral hump, about 74-110 setae on ventral surface. Gills absent. Other abdominal structures as in Goeracea. Larval case (Fig. 19) of small rock fragments, lacking the lateral rows of larger pieces characteristic of species in Goeracea; posterior opening restricted with silk, leaving only a small opening dorsal to centre as in Goeracea. Length of larval case to 6.4 mm. PUPA. Only G. betteni available for study; structure essentially as in Goeracea. Anal processes taper to a more slender point and curl dorsad. Closure membranes of pupal case similar to Goeracea but anterior meshes somewhat more open and sieve-like. ADULT. Apart from genitalic characters, this genus can be distinguished from Goeracea and Lepania by the swollen segments of the maxillary palpi of the male. Length of forewing 4.8-6.4 mm. Colour light brown. Head (Fig. 24b) with dorsal, posterolateral warts broader than in Goeracea, long axis of warts nearly parallel to that of the body; one pair of dorsal anteromesal warts; ocelli lacking; maxillary palpi of male (Fig. 24a) with middle and terminal segments swollen, terminal, semi-membranous. Thorax similar to that illustrated for Lepania (Fig. 22). Forewings without scales; frenulum of 4-5 stout setae as in Goeracea. Venation similar to Goeracea, and as illustrated by Ross (1944, fig. 874). Spurs 2, 4, 4. Male genitalia (Ross, 1944, fig. 878; 1962, fig. 3). Distinctive in the structure of segment x, where the praeanal appendages lie close together along the middorsal line, overlying the membranous dorsum of the segment; other appendages of segment X apparently lacking. Claspers much as in Goeracea, basal segment lacking the prominent ventromesal process of Goeracea; terminal segment finger-like, with stout peg-like teeth on mesal surface. Female genitalia (Fig. 34). Distinguished by the broadly-rounded median lobe separating the posterolateral sclerites of segment 1x. Dorsum of segment Ix evidently separated from x by a membranous area. 16 HABITAT. Larvae of both species occur in small, cold, spring runs. Key to species of Goerita 2. Median ridge of pronotum evenly confluent laterally (CEC gel (5) Ge achat Once Or Ae eee en ne SS ...... G. betteni Median ridge of pronotum with a prominent lateral excavation, posterior surface of ridge also EXC Anace (ED ZO ern Bee ee en Rh iy feo ls ue. G. semata ee Walmer se oy oe. gone ew. ee ee ee 2 2 a MR lps yh 2 ieee 4 Female ... te alee, ts BM scl ep et | A 5 4. Segment x with mesal margins of praeanal append- ages straight and approximate along the middorsal line for most of their length (Ross, 1944, fig. 878) 0.00000... G. semata Segment xX with mesal margins of praeanal append- ages curved and approximate only near the apex (Ross; 1962 shige Si) us eee RS - G. betteni 5. Median plate of venter 1x broadly rounded apically (Fis. 84) ee. Ree eee AS .......... G. semata Female of G. betteni not known. Goerita semata Ross Goerita semata Ross. 193879, 1/2, fig. 113. Goerita semata. Ross, 1944, p. 257, figs. 874, 875, 878. Goerita semata. Flint, 1960, p. 39, fig. 20. No firm association has been established between larvae and adults of this species. But collections of unique larvae, apparently congeneric with larvae of G. betteni and taken in the general vicinity of the type locality of G. semata, support the circumstantial association proposed by Flint (1960). LARVA (Fig. 20). Length of final-instar larva referred to above: 4.8-6.1 mm (25); length of larval case 5.8-6.4 mm (5). Abdominal segments (final instar): I, setae between median hump and each lateral hump 41-52 (5); setae on ventral surface 94-112 (5); 1x, setae on dorsal selerite 38-49 (5). PUPA. None available. iF Y CP e Yj WY ies x it PS Vt , Faw WD i VS ganguro Wins oat ae Cite 0 in ; "! ‘ ily aa £4 J) ik to, 190 <> SOA ND lh, r ih ANY Ir Ce OD 3, nll!’ @RGOar \\P EY \ 7, \\\\\\ Al Y Yy GY Yj Gy, va b J EN ADULT. Length of forewing: male, 4.8-5.4 mm (4); female 6.4 mm (4d¢d, [Oo stype series). BIOLOGY. Larvae occur on the undersides of rocks. In contrast to G. betteni, larvae of this species use rock fragments of more uniform size in con- structing cases. Other observations are provided by Flint (1960). DISTRIBUTION. This species is known only from localities in mountains of the southeastern United States. Figs. 17-19 Goerita betteni larva. 17. Larva: a, lateral view, X24; b, ninth abdominal segment and anal prolegs, dorsal view, X53.5. 18. Head and thorax, dorsal view, X48. 19. Larval case: a, lateral view, X17: b, posterior end, X17. Fig. 20 Goerita semata larva, head and thorax, dorsal view, X59. 19 MATERIAL STUDIED. NORTH CAROLINA: Mount Mitchell, Camp Alice Road, 3 Sept. 1959, G.B.W. and O.S. Flint, 5 larvae. TENNESSEE: Spring stream at Indian Gap, Great Smoky Mountains National Park, 13 Sept. 1958, G.B.W. and O.S. Flint, many larvae; small stream in Indian Gap, Great Smoky Mountains National Park, elevation approximately 5200 ft, 20 May 1970 (700358) G.B.W. and T. Yamamoto, many larvae. Goerita betteni Ross Goerita betteni Ross, 1962, p. 132, fig. 3. Dr. S.W. Edwards has kindly made available to me some of the material upon which his records of G. semata from Wellington Mills, Tennessee were based (Edwards, 1966). These specimens, all males, are G. betteni. It is noteworthy that his published records from this locality indicate occur- rence of adults in late April and in September. LARVA (Figs. 17-19). Length of final-instar larva 4.4-6.1 mm (25); length of larval case 5.6-6.4 mm (5). Abdominal segments (final instar): I, setae between median hump and each lateral hump 20-28 (5); setae on ventral surface 74-88 (5); Ix, setae on dorsal sclerite 31-42 (5). PUPA. Length of pupa c. 4.6 mm (1 specimen, damaged). ADULT (Figs. 24, 34). Length of forewing 4.6-5.6 mm (96d). Males (Fig. 24) differ from those of G. semata in having the middle segment of the maxillary palp less quadrate (cf. Ross, 1944, fig. 113), the posterodorsal warts of the head broader, and the eyes larger (in dorsal view the maximum width of the eye in G. betteni is approximately equal to one-half of the interocular width but in G. semata to approximately one-quarter of the interocular width). BIOLOGY. Larvae were located primarily on vertical rock faces in a thin layer of flowing spring water. Based upon the single population that we sampled, the case-making behaviour of G. betteni differs from that of G. semata in the selection of larger rock fragments for the sides of the case (Fig. 19) than for the dorsal and ventral surfaces. This behaviour suggests a modification of that in Goeracea, in which larvae fasten much larger stones along each side of the case (Fig. 12). DISTRIBUTION. This species is now known from West Virginia, Tennessee, and perhaps New York (Ross, 1962). MATERIAL STUDIED. TENNESSEE. Franklin Co.: Hillside spring, Welling- ton Mills, 4 mi W on Route 50, 1.5 mi N Winchester, 14-15 May 1970 (700337) G.B.W. and T. Yamamoto, many larvae, 1 pupa, 2dd; same locality, 13 Sept. 1955, S.W. Edwards, 7é6, and 399 (dates. lacking), S.W. Edwards collection. 20 Systematic Considerations for Lepania cascada Assignment of Lepania cascada to the Apataniinae (Moropsychini) was based primarily on characters of the genitalia (Schmid, 1968a), and in the absence of any knowledge of the immature stages of either group. Now available from Dr. Schmid’s field work in India are larvae and pupae of two species of Moropsyche, the largest genus of the Moropsychini. Analysis of this material (Botosaneanu, 1968) indicated that larval and pupal characters in at least the genus Moropsyche are largely consistent with those in the subfamily Apataniinae. Evidence presented here from larval characters in Lepania reveals that the relationship between Lepania and Moropsyche is so distant that the two genera cannot be considered as members of the same subfamily. Larvae of L. cascada clearly have the general facies of a member of the limnephilid subfamily Goerinae. This larval facies is so characteristic of the Goerinae, and unique to that group of all the Trichoptera, that its presence in Lepania must be regarded as potentially significant. Comparative analysis of characters of all stages of L. cascada which follows should either substantiate this relationship or, in failing to do so, indicate that similarity in larval facies is probably the result of convergent evolution, the true genetic affinities of the genus lying elsewhere. 1. Larva. Students of the Trichoptera are agreed on a close relationship between the subfamilies Apataniinae and Goerinae. Some of the best evidence for this is derived from larvae, and most larval characters shared by the two groups represent what evidently are derived conditions of ancestral limnephilid characters (Wiggins, in press). Larval characters unique to the Goerinae, and therefore distinguishing them from the Apataniinae, are (a) pronotum (Fig. 10a) enlarged dorsally, forming a sharp, lateral carina, ventral to which the pronotum is thickened; (b) mesonotum (Fig. 18) subdivided into two or three pairs of plates, with a prominent transverse ridge forming an axis on which the whole anterior part of the larva is canted ventrad (Fig. 10a); (c) mesopleuron enlarged, mesepisternum extended into a long, stout, pointed process (Fig. 20). Com- parison of the larva of Lepania cascada (Figs. 1 and 2) with these characters, unique to the Goerinae, shows complete agreement. , Additional characters in the larva of L. cascada may indicate a goerine relationship: (a) On segment x, ventrolateral to the basal tuft, is a small sclerite (Fig. 1c). In Goeracea and Goerita the two large setae of the basal tuft are encircled by a small sclerite (Figs. 10c, 17b). The similarity between the sclerites in these genera suggests that they may be homologous. This sclerite appears to be a homologue of the dorsal plate (sensu Ross, 1956, p. 13) of the Rhyacophiloidea, a structure absent in most groups of the Limnephi- loidea. The presence of a sclerite ventral to the lateral sclerite and basal tuft of the anal prolegs is not known in the Apataniinae. (b) The structure of the larval case in L. cascada (Fig. 4a) shows that the larva places smaller stones ventrally and larger ones laterally and dorsally. Positioning of stones in this way is an unusual feature of case- 21 Figs. 21-23 Lepania cascada. 21. Wings: a, wings of right side, X11; b, detail of wing- coupling structures. 22. Head and thorax of male, dorsal view, X40. 23. Head of male, facial view, X62. building in the Trichoptera, but finds a parallel in behaviour in Goerita (Fig. 19a). 2. Pupa. Among the pupal characters of L. cascada, the smoothly curving, laterally dentate anal processes (Fig. 7) are more similar in shape to those of the Moropsychini (Botosaneanu, 1968) than to those of the Goerinae. In the Moropsychini, however, these processes bear two pairs of stout setae near the apex, as compared with three pairs in Lepania. The only goerine genera available to me as pupae, Goera, Goeracea (Fig. 14), and Goerita, are so different in this respect that no clear goerine character emerges for the anal processes, although the anal processes do bear three pairs of apical setae in Goeracea and Goerita, as in Lepania, but none in Goera. The primi- tive condition in the Limnephilidae appears to be three setae near the apex of each process, with two characteristic for the Apataniinae. Ze Fig. 24 Goerita betteni, head of male: a, facial view, X58; b, dorsal view, X62.5. The pupal case in Moropsyche (Botosaneanu, 1968, fig. 127) appears to have the characteristic apataniine hood over the anterior opening. The pupal case in Lepania does not have such a hood, nor do pupal cases in Goera, Goeracea, or Goerita. On the other hand, the silken membranes closing each end of the pupal case in Lepania appear to be similar to those in the Moro- psychini (Botosaneanu, op. cit., fig. 128). A pair of very long, hooked setae on the mid part of the frontal area of L. cascada (Fig. 5) appears to be distinctive for this genus alone. Other pupal characters seem neither to be distinctive nor to suggest relationship with any particular group. 3. Adult. Adults in Lepania possess the basic diagnostic characters required for assignment to the Limnephilidae (s.s. excluding Goerinae) — ocelli, max- illary palpi three-segmented in the male and five-segmented in the female. Other characters of the adults appear to be so concordant that this assign- ment has not been questioned previously. But several characters in Lepania adults are concordant with those found in at least some species of the Goerinae. I do not have sufficient material of the world fauna of the Goerinae to carry this analysis further now or to assess primitive and derived char- acters. But until that should be possible, the following evidence demonstrates that numerous characters are possessed in common by Lepania and the Goerinae, including several that probably are derived. (a) The three-segmented maxillary palpi of the male in Lepania (Fig. 23) are short and curve vertically in front of the face as in Goeracea. In the Limnephilidae (s.s.) the maxillary palpi are generally longer and do not curve vertically in front of the face. Pe 272] preape: == L. cascada -int. br., seg. X-——- ext. br., seg. X—-— acc. app.-— —-- 26 G. archaon (b) Only one pair of anterior warts occurs posterior to the median ocellus in Lepania (Fig. 22), as in Goeracea, Goerita, and Goera. The mesoscutel- lum in Lepania bears a single, ovoid, median wart (Fig. 22). In the Apata- niinae, as far as I have been able to ascertain, there are two pairs of warts posterior to the median ocellus, and the mesoscutellum bears a pair of small warts. Although the arrangement of these warts in Lepania is similar to some other limnephilids, this discordance with the Apataniinae does not support assignment of Lepania to that subfamily. (c) In the hind wing in Lepania, R, is atrophied or fused with Sc (Fig. 21), as it is in some other species of the Goerinae, e.g., Silo nigricornis (Mosely, 1939, fig. 227). This vein is characteristically strongly developed throughout the Limnephilidae (s. s.). (d) The wing-coupling mechanism in L. cascada (Fig. 21) consists of a row of stout, hooked setae on the costal margin of the hind wing. These engage with a series of stout hairs arising along the midanal margin of the forewing. The mechanism is essentially the same in Goera, although a frenulum also occurs in the latter. The wing-coupling mechanism in the Limnephilidae consists basically of a row of short bristles on the costal margin of the hind wing, frequently with a patch of longer setae near the base comprising the frenulum, but appears to lack a fringe of stout hairs on the forewing. The frenulum in the Apataniinae is further characterized by comprising three stout, apically clavate hairs (Schmid, 1953, fig. 18). (e) The male genitalia of Lepania cascada (Fig. 25) were considered (Schmid, 1968a) to provide the best evidence for relationship with the Apataniinae (Moropsychini). Several characters are, however, shared with the Goerinae. In L. cascada, a membranous area on the sternum of segment Ix (Fig. 25c), bordered laterally by linear thickenings, was regarded by Schmid (1968a) as an important and exclusive character of Lepania. In Goeracea genota (Fig. 29c) two similar linear thickenings appear on the ninth ster- num, the area between them grading from lightly sclerotized anteriorly to membranous posteriorly. Segment x bears, in addition to the praeanal appendages, three pairs of appendages in Lepania (Figs. 25, 26) and two in the Moropsychini, inter- preted in the latter as the external and internal branches of segment x (Schmid, 1968b). Two of the pairs of appendages on segment x of L. cascada could also be so interpreted, but the third pair, here designated acces- sory appendages of segment x, was regarded by Schmid (1968a) as an important and exclusive character of the genus Lepania. Segment x of Goera archaon (Fig. 26) also bears three sets of appendages in addition to the Figs. 25-27 Male genitalia. 25. Lepania cascada: a, lateral view, X100; b, dorsal view, X100; c, ventral view, X100; d, phallus, ventral and lateral views, X88. 26. Tenth segments of L. cascada and Goera archaon, dorsal view; acc. app. accessory appendages; ext. br., seg. X — external branch, segment X; int. br., seg. X — internal branch, segment X; pr. app. — praeanal append- age. 27. G. archaon: a, lateral view, X80; b, dorsal view, X80; c, phallus, lateral view, X87. 25 pr. app.,seg. X oregona Figs. 28-29 Male genitalia. 28. Goeracea oregona: a, lateral view, X100; b, dorsal view, X100; c, ventral view, X100; d, phallus, ventral and lateral views, X 100; pr. app., seg. X — praeanal appendage, segment X. 29. Goeracea genota: a, dorsal view, X102; b, ventral view, X102. ger ie sclerite, seg. VIII cascada lateral sclerite, Figs. 30-34 Female genitalia. 30. Lepania cascada: a, lateral view, X72; b, ventral view, X72. 31. Goera stylata, female genitalia, ventral view, X54. 32. Goeracea oregona: a, lateral view, X57; b, ventral view, X57; c, caudal view, X57. 33. Goeracea genota: a ventral view, X48.5; b, caudal view, X48.5. 34. Goerita semata: a, lateral view, X67; b, ventral view, X67. yf praeanal pair, and the position and shape of the accessory appendages in both species are remarkably similar. The praeanal appendages in L. cascada are unusually large, but in Goeracea genota (Fig. 29) and in Larcasia spp. (Schmid, 1965) they are also large. G. genota provides an especially instructive comparison because in this species the appendages are bifid, as in Lepania. (f) The female genitalia of Lepania cascada were thought to offer fewer characters than those of the male to support the relationship with the Moro- psychini (Schmid, 1968a), although segment x in Lepania was said to be typically apataniine. My interpretation of the tenth segment in the Apata- niinae is that it is fused with rx to form a prominent semi-cylinder surround- ing the anal opening, and that the supragenital plate is fused across the ventrolateral edges of this semi-cylinder. Segment x in L. cascada (Fig. 30) is not of this precise type and lacks the heavily sclerotized, semi-cylinder structure. My interpretation is that segment x in Lepania shows primitive features in common with several genera of the Limnephilidae (Imania, Moselyana, and Goera) but that the Apataniinae have become specialized in a unique way, different from Lepania. The ventromedian lobe of the terminal abdominal segment of apataniine females was interpreted (Schmid, 1953, 1954, 1955) as being derived from segment vill. The presence of an apparently complete sternum to segment vill in Lepania cascada (Fig. 30), and a partial sternum to segment vil in Goeracea spp. (Figs. 32, 33) suggests that in these species the ventromedian apical lobe could be interpreted as part of the venter of segment Ix. In any event, venter vill of L. cascada is more similar to that of Goeracea spp. than to any of the Apataniinae, as demonstrated by the darkened sclerites lateral to the median plate (cf. Figs. 30, 32). Comparison of the posteroventral margin of the apical abdominal segment in L. cascada (Fig. 30b) with that in Moropsyche spp. (Schmid, 1968b) shows little basic similarity between the two. Comparison of L. cascada with Goera stylata (Fig. 31) shows, however, a close concordance between these two species in the structure of the posteroventral margin of the apical seg- ment. Here again affinity of Lepania with the Goerinae is indicated. In limne- philid genera that have some general similarity to the Apataniinae, such as Imania and Moselyana, a prominent median lobe terminates the apical segment, which is usually transversely striated and semi-membranous (Schmid, 1968a; Wiggins, in press). The same is true of Moropsyche (Schmid, 1968b). In the Goerinae the median lobe appears to be represented by a simple, triangular plate, and this is precisely what occurs in Lepania. The tergum of segment Ix in L. cascada females is not distinct from that of segment x (Fig. 30a) and may either be fused with the latter beyond recognition or be absent altogether. In the Apataniinae the fusion of seg- ments Ix and x is usually obvious through the presence of a ridge or promi- nent suture. In Goeracea, on the other hand, a small, separate sclerite appears (Fig. 32a) in the position of the tergum of segment Ix, suggesting that it is separate from x and disappearing. The condition of the dorsal parts of segments Ix and x in Lepania is clearly more similar to that in Goeracea than to the Apataniinae. 28 | : | | | Conclusions 1. CLASSIFICATION. The foregoing evidence shows that the general resem- blance of the larva of Lepania cascada to the Goerinae is fully borne out by detailed examination of larval characters. Among the pupal characters of Lepania, some are similar to those of the Moropsychini, others to those of the ancestral limnephilid condition, and others unique. But assessment of the significance of pupal characters is complicated by the fact that many are so diverse that it is difficult to derive a consensus of characters, and also difficult to assess ancestral and derived conditions. Characters of adults, both genitalic and non-genitalic, yield a surprising amount of evidence for affinity of Lepania with the Goerinae. Available evidence shows that the genus Lepania cannot be assigned to the Apataniinae, as previously proposed, but rather that this genus has more in common with the limnephilid subfamily Goerinae. I propose that Lepania be assigned to this group. Retention of certain primitive characters in this genus, however, requires creation of a new and distinct tribe. Lepaniini new tribe type genus Lepania Ross, 1941, p. 102 DIAGNOSIS. The tribe Lepaniini can be distinguished from other taxa of the Goerinae in the larval stage by the presence of separate teeth on the man- dibles (Fig. 3) and by the basal seta on the tarsal claw, which extends far short of the tip of the claw (Fig. 1b). In all other larvae of the Goerinae now known, the mandibles have a single cutting edge (Fig. llc), and the basal seta of each tarsal claw extends nearly to the tip of the claw (Fig. 10b). Pupae can be distinguished by the pair of long setae in the middle of the frontal area (Fig. 6) and by the curved, dentate anal processes (Fig. 7). In the adult stage, the Lepaniini can be distinguished from most other mem- bers of the Goerinae by the presence of ocelli (Fig. 22). In the genus Goereilla, recently described by Denning (1971) and assigned to the Goeridae, the single species G. baumanni Denning has ocelli. This species, from Montana, is known only from males, and confirmation of its systematic position awaits discovery of females and larvae. Dr. Denning loaned me a male of G. baumanni for study, and this sex, at least, shows little similarity to L. cascada. The maxillary palpi are short overall; the basal segment is short and sub-spherical, the middle and terminal segments elon- gate and approximately equal in length. R, in the hind wing is fully devel- oped; the hind wing has a frenulum of several long, straight setae, and a row of shorter, straight setae along the costal margin. The genitalia (Denning, 1971, fig. 5) show little resemblance to those of L. cascada. Two pairs of warts posterior to the median ocellus also distinguish adults of Goereilla from Lepania, in which there is only a single pair. The recognition of Lepania as an annectent form between the goerine line and the Limnephilidae (s. s.) introduces a new dimension to the subject of the two versions of the classification of this line: subfamily Goerinae of the Limnephilidae (Nielsen, 1943; Flint, 1960); or the Goeridae as a distinct 29 family (Schmid, 1955; Ross, 1967). There is no disagreement that the goerid and limnephilid lines are closely related. Details of classification of this sort are, of course, ultimately subjective, but the evidence presented here for Lepania does narrow the taxonomic gap between the goerid and limnephilid lines and strengthens the view that the gap is not sufficiently broad to warrant recognition of two distinct families. From the standpoint of practical taxonomy, a general diagnosis between the two groups now becomes increasingly difficult, and this ambiguity seems less appropriate to the level of family than subfamily. An assessment of the characters of the adults made by Schmid (1955, p. 16) is perhaps more relevant now than it was at that time: “La famille des Goérides est incontestablement la plus proche parente des Limnophilides. Certains de ces représentants mémes pourraient étre considérés comme ap- partenant a cette famille, n’étaient la réduction des palpes maxillaires et Pabsence d’ocelles.” Perhaps the ultimate test has already been made, for in assigning adults of Lepania cascada to the Limnephilidae (s. s.) for 30 years, workers have demonstrated how narrow is the gap between that family and the Goeridae. 2. PHYLOGENY. (a) The combination of characters found in Lepania has interesting implications for the phylogeny of the Goerinae and related genera of the Limnephilidae. Three characters of Lepania are particularly relevant: (i) presence of ocelli in the adult (Fig. 22); (ii) larval mandibles with well- developed teeth (Fig. 3); (iii) basal seta of tarsal claws of larva much shorter than the claw (Fig. 1b). These all appear to be primitive characters that likely occurred in the ancestral limnephilids (Wiggins, in press). All three are represented in a derived condition in the typical Goerinae: (1) ocelli lost; (ii) mandibles with separate teeth united to form a single edge (Fig. llc); (iii) basal seta of larval tarsal claws extending almost to the tip of the claw (Fig. 10b). Assuming that the condition of these characters in L. cascada does not represent a reversion to the primitive type from a previously derived condi- tion, then Lepania must be regarded as the most primitive living taxon of the Goerinae now known, allowing for the possible addition of the genus Goereilla to this category when its relationships are clarified (see 1 above). Evidence for this primitive position from three apparently independent characters of adult and larva adds strength to the interpretation. The close relationship postulated between the Limnephilidae (s. s.) and the Goeridae (auct.) has been based on the hypothesis that the annectent forms possessed ocelli in the adult (Ross, 1967). Lepania, as characterized here, may be regarded, then, as demonstrating the validity of this hypothesis and is therefore a genus of unusual phylogenetic significance. If L. cascada were an extinct species, perfectly preserved in a piece of fossil amber, and by some remarkable series of events clearly associated through larva, pupa, male, and female, one could suspect that its position as an annectent form between the ancestral limnephilid stock and the goerine line of evolution would be much enhanced. (b) Structural modifications in the larval head and thorax of the Goerinae 30 are, as far as is now known, unique in the Trichoptera. Observation of living larvae shows that these structures are integrated to make a very tight closure to the anterior end of the larval case. Lepania has typical goerine character- istics: elongate mesepisternum; angulation of the thorax and head across the transverse dorsal ridge of the mesonotum; and subdivision of the mesono- tal plates into separate sclerites. But the head is not recessed into the prono- tum to help form the closure of the case opening, as in Goera, or curled sharply ventrad beneath the pronotum as in Goeracea genota. In the Goerinae, it seems likely that the concavities in which the antennae lie are related to the need for compaction of the head into minimal space within the case. Closure of the case by the withdrawn larva seems to have evolved to a level of precision in the higher Goerinae that is not seen in any other group, yet it is not yet clear why this should be so. Most goerine larvae live on rocks in rapid streams, and their need for protection from intruding predators seems little different from the needs of similar caddis larvae of their community — e.g., Neophylax spp. or Pycnopsyche spp. in North American waters. An interesting coincidence is that larvae of at least some of the higher Goerinae (I have not seen larvae of extra-North American genera) also restrict with silk the opening in the posterior end of their cases to a much smaller hole (e.g. Fig. 12b) than is usual in genera of the Limnephilidae (Sa53)) The ability of the larva of Lepania cascada to close the anterior opening of its case with head and thorax seems not as well perfected as in the higher goerines; and the opening in the posterior end of the case is not reduced as much (Fig. 4b). Whatever the evolutionary stimulus causing this behaviour in the higher goerines may have been, the Lepaniini appear to represent an earlier, transitional stage. Acknowledgments Field studies during which specimens and observations were obtained for this paper were supported initially by the National Science Foundation (G22135), and later by the National Research Council of Canada (A5707). This support is gratefully acknowledged. Several colleagues kindly gave assistance of many kinds for my field studies: Dr. N. H. Anderson, Depart- ment of Entomology, Oregon State University; Mr. S. G. Jewett, Jr., Portland, Oregon; Dr. A. L. Sheldon, Sagehen Creek Research Project, University of California (now at the Department of Zoology, University of Montana). For assistance with field operations, I am indebted to Messrs T. Yamamoto, I. M. Smith, and R. S. Scott. Mr. Yamamoto also assisted in the organization of systematic data. For the loan of specimens from the collection of the Illinois Natural History Survey I am indebted to Dr. John D. Unzicker. Dr. D. G. Denning, Moraga, California, kindly loaned specimens from his col- lection, as did Dr. S. D. Smith, Central Washington State College, and Dr. S. W. Edwards, Southwest Texas State University. The illustrations in the paper were prepared by Mr. Anker Odum, scientific illustrator, Department of Entomology and Invertebrate Zoology, ROM. I am indebted to Dr. F. Schmid, Ottawa, for critical comments on the manuscript. aT Literature Cited BOTOSANEANU, L. 1968 Les jeunes stades des Moropsychini: Moropsyche vanegudha n.sp. et krichnaruna n.sp. In Schmid, F. La sous-famille des Apataniines en Inde (Trichoptera, Limnophilidae). Can. Ent., vol. 100, no. 12, pp. 1269-1277. DENNING, D. G. 1968 New and interesting North American Trichoptera. Pan-Pacif. Ent., vol. 44, no. 1, pp. 17-26. 1971 A new genus and new species of Trichoptera. Pan-Pacif. Ent., vol. 47, no, 3,4pp) 202-210. EDWARDS, S. W. 1966 An annotated list of the Trichoptera of middle and west Tennessee. J. Tenn. Acad: Sci., vol. 41, no. 4, pp. 116-128. FLINT .O; S.,- JR: 1960 Taxonomy and biology of nearctic limnephilid larvae (Trichoptera), with special reference to species in eastern United States. Entomologica Am., n.s., vol. 40, pp. 1-117. MOSELY, M. E. 1939 The British caddis flies (Trichoptera); a collector’s handbook. London, George Routledge. 320 pp. NIELSEN, A. 1943 Trichopterologische Notizen. Vidensk. Meddr. Dansk Naturh. Foren., vol. 107, pp. 105-120: ROSS, H. H. ° 1938 Descriptions of nearctic caddis flies (Trichoptera), with special reference to the Illinois species. Bull. Ill. St. Nat. Hist. Surv., vol. 21, art. 4, pp. 101-183. 1941 Descriptions and records of North American Trichoptera. Trans. Am. Ent. Soc., vol. 67, pp. 35-126. 1944 The caddis flies, or Trichoptera, of Illinois. Bull. Ill. St. Nat. Hist. Surv., Vol! (23; "art *4 ar a Pe ‘ ‘ > i 3 7 ee es 7 ——. ~ 4 ‘ , : “~ me fi oo ‘ae - A se eo ae ea 1) . ie | . f > as ap ie wis. .' 24, oy! 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